Behavioural thermoregulation in cold-water freshwater fish: Innate resilience to climate warming?

Behavioural thermoregulation enables ectotherms to access habitats providing conditions within their temperature optima, especially in periods of extreme thermal conditions, through adjustments to their behaviours that provide a "whole-body" response to temperature changes. Although freshwater fish have been detected as moving in response to temperature changes to access habitats that provide their thermal optima, there is a lack of integrative studies synthesising the extent to which this is driven by behaviour across different species and spatial scales. A quantitative global synthesis of behavioural thermoregulation in freshwater fish revealed that across 77 studies, behavioural thermoregulatory movements by fish were detected both vertically and horizontally, and from warm to cool waters and, occasionally, the converse. When fish moved from warm to cooler habitats, the extent of the temperature difference between these habitats decreased with increasing latitude, with juvenile and non-migratory fishes tolerating greater temperature differences than adult and anadromous individuals. With most studies focused on assessing movements of cold-water salmonids during summer periods, there remains an outstanding need for work on climatically vulnerable, non-salmonid fishes to understand how these innate thermoregulatory behaviours could facilitate population persistence in warming conditions.

Marine heatwaves impair the thermal refugia potential of marginal reefs in the northern South China Sea.

Frequent marine heatwaves (MHWs), concurrent with climate warming, threaten global low-latitude, pristine coral reefs, leading to growing interest in identifying marginal coral reefs (relatively high-latitude and/or turbid reef environments) that can serve as thermal refugia from mass coral bleaching. However, the thermal refugia potential of marginal reefs remains controversial. We evaluated the thermal refugia potential of inshore reefs in the northern South China Sea (nSCS), a globally typical marginal reef system, by characterizing the long-term trend of MHW intensity and frequency and assessing thermal stress during a mass bleaching event in summer 2020. An unprecedented peak intensity of around 20 °C-weeks of cumulative heat stress, associated with a prolonged anomalous western Pacific subtropical high (WPSH) and weakened monsoon activity, induced record-breaking bleaching. The geographical variability of bleaching was strongly related to the extent of heat exposure and satellite-derived temperature anomalies. Under ongoing global warming, the frequency and intensity of MHWs over nSCS coral habitats show a markedly increasing trend, especially during the last decade. Intense MHWs and coral bleaching have already occurred throughout all El Niño-Southern Oscillation (ENSO) phases (e.g., 2010, 2015, and 2020). Climate change has pushed marginal coral reefs to or beyond the limits of their resilience, and frequent MHW events have amplified the increasing risk of thermal stress. There are no long-term thermal refugia for marginal reefs in the nSCS.

Individual behaviour and resource use of thermally stressed brook trout Salvelinus fontinalis portend the conservation potential of thermal refugia.

Individual aggression and thermal refuge use were monitored in brook trout Salvelinus fontinalis in a controlled laboratory to determine how fish size and personality influence time spent in forage and thermal habitat patches during periods of thermal stress. On average, larger and more exploratory fish initiated more aggressive interactions and across all fish there was decreased aggression at warmer temperatures. Individual personality did not explain changes in aggression or habitat use with increased temperature; however, larger individuals initiated comparatively fewer aggressive interactions at warmer temperatures. Occupancy of forage patches generally declined as ambient stream temperatures approached critical maximum and fish increased thermal refuge use, with a steeper decline in forage patch occupancy observed in larger fish. These findings suggest that larger individuals may be more vulnerable to stream temperature rise. Importantly, even at thermally stressful temperatures, all fish periodically left the thermal refuge to forage. This indicates that the success of refugia at increasing population survival during periods of stream temperature rise may depend on the location of thermal refugia relative to forage locations within the larger habitat mosaic. These results provide insights into the potential for thermal refugia to improve population survival and can be used to inform predictions of population vulnerability to climate change.

Inferring watershed hydraulics and cold-water habitat persistence using multi-year air and stream temperature signals.

Streams strongly influenced by groundwater discharge may serve as "climate refugia" for sensitive species in regions of increasingly marginal thermal conditions. The main goal of this study is to develop paired air and stream water annual temperature signal analysis techniques to elucidate the relative groundwater contribution to stream water and the effective groundwater flowpath depth. Groundwater discharge to streams attenuates surface water temperature signals, and this attenuation can be diagnostic of groundwater gaining systems. Additionally, discharge from shallow groundwater flowpaths can theoretically transfer lagged annual temperature signals from aquifer to stream water. Here we explore this concept using multi-year temperature records from 120 stream sites located across 18 mountain watersheds of Shenandoah National Park, VA, USA and a coastal watershed in Massachusetts, USA. Both areas constitute important cold-water habitat for native brook trout (Salvelinus fontinalis). Observed annual temperature signals indicate a dominance of shallow groundwater discharge to streams in the National Park, in contrast to the coastal watershed that has strong, apparently deeper, groundwater influence. The average phase lag from air to stream signals in Shenandoah National Park is 11 d; however, extended lags of approximately 1 month were observed in a subset of streams. In contrast, the coastal stream has pronounced attenuation of annual temperature signals without notable phase lag. To better understand these observed differences in signal characteristics, analytical and numerical models are used to quantify mixing of the annual temperature signals of surface and groundwater. Simulations using a total heat budget numerical model indicate groundwater-induced annual temperature signal phase lags are likely to show greater downstream propagation than the related signal amplitude attenuation. The measurement of multi-seasonal paired air and water temperatures offers great promise toward understanding catchment processes and informing current cold-water habitat management at ecologically-relevant scales.

Badger setts provide thermal refugia, buffering changeable surface weather conditions.

Den use can be crucial in buffering environmental conditions and especially to provide an insulated environment for raising altricial young. Through Sept-Dec 2016 we monitored temperature and humidity at 11 badger setts (burrow systems), using thermal probes inserted over 4-13 sett entrances to a depth of ca. 2 m, supplemented by continuous daily logging at one entrance per sett. Setts were cooler than exterior conditions Sept-Oct, and warmer than exterior conditions Nov-Dec. Setts cooled down when badgers left them to forage by night, and warmed up when badgers occupied them by day. Soil type and aspect also influenced sett temperature. Sett temperature did not affect the weight or body-condition of either adults or maturing cubs in autumn. However, cubs born into setts that were relatively warmer through the preceding autumn-winter were heavier in the following spring than contemporaries born in cooler setts (badgers exhibit delayed implantation), and so warmer setts might benefit early cub growth. We posit that sett quality may be important in providing badgers with a stable thermal refuge from variable weather conditions. More broadly, den use may buffer climate change effects for many fossorial carnivore species.

Can brook trout survive climate change in large rivers? If it rains.

We provide an assessment of thermal characteristics and climate change vulnerability for brook trout (Salvelinus fontinalis) habitats in the upper Shavers Fork sub-watershed, West Virginia. Spatial and temporal (2001-2015) variability in observed summer (6/1-8/31) stream temperatures was quantified in 23 (9 tributary, 14 main-stem) reaches. We developed a mixed effects model to predict site-specific mean daily stream temperature from air temperature and discharge and coupled this model with a hydrologic model to predict future (2016-2100) changes in stream temperature under low (RCP 4.5) and high (RCP 8.5) emissions scenarios. Observed mean daily stream temperature exceeded the 21°C brook trout physiological threshold in all but one main-stem site, and 3 sites exceeded proposed thermal limits for either 63- and 7-day mean stream temperature. We modeled mean daily stream temperature with a high degree of certainty (R2=0.93; RMSE=0.76°C). Predicted increases in mean daily stream temperature in main-stem and tributary reaches ranged from 0.2°C (RCP 4.5) to 1.2°C (RCP 8.5). Between 2091 and 2100, the average number of days with mean daily stream temperature>21°C increased within main-stem sites under the RCP 4.5 (0-1.2days) and 8.5 (0-13) scenarios; however, no site is expected to exceed 63- or 7-day thermal limits. During the warmest 10years, ≥5 main-stem sites exceeded the 63- or 7-day thermal tolerance limits under both climate emissions scenarios. Years with the greatest increases in stream temperature were characterized by low mean daily discharge. Main-stem reaches below major tributaries never exceed thermal limits, despite neighboring reaches having among the highest observed and predicted stream temperatures. Persistence of thermal refugia within upper Shavers Fork would enable persistence of metapopulation structure and life history processes. However, this will only be possible if projected increases in discharge are realized and offset expected increases in air temperature.

Thermal variability alters the impact of climate warming on consumer-resource systems.

Thermal variation through space and time are prominent features of ecosystems that influence processes at multiple levels of biological organization. Yet, it remains unclear how populations embedded within biological communities will respond to climate warming in thermally variable environments, particularly as climate change alters existing patterns of thermal spatial and temporal variability. As environmental temperatures increase above historical ranges, organisms may increasingly rely on extreme habitats to effectively thermoregulate. Such locations desirable in their thermal attributes (e.g., thermal refugia) are often suboptimal for resource acquisition (e.g., underground tunnels). Thus, via the expected increase in both mean temperatures and diel thermal variation, climate warming may heighten the trade-off for consumers between behaviors maximizing thermal performance and those maximizing resource acquisition. Here, we integrate behavioral, physiological, and trophic ecology to provide a general framework for understanding how temporal thermal variation, mediated by access to a thermal refugium, alters the response of consumer-resource systems to warming. We use this framework to predict how temporal variation and access to thermal refugia affect the persistence of consumers and resources during climate warming, how the quality of thermal refugia impact consumer-resource systems, and how consumer-resource systems with fast vs. slow ecological dynamics respond to warming. Our results show that the spatial thermal variability provided by refugia can elevate consumer biomass at warmer temperatures despite reducing the fraction of time consumers spend foraging, that temporal variability detrimentally impacts consumers at high environmental temperatures, and that consumer-resource systems with fast ecological dynamics are most vulnerable to climate warming. Thus, incorporating both estimates of thermal variability and species interactions may be necessary to accurately predict how populations respond to warming.

Loss of thermal refugia near equatorial range limits.

This study examines the importance of thermal refugia along the majority of the geographical range of a key intertidal species (Patella vulgata Linnaeus, 1758) on the Atlantic coast of Europe. We asked whether differences between sun-exposed and shaded microhabitats were responsible for differences in physiological stress and ecological performance and examined the availability of refugia near equatorial range limits. Thermal differences between sun-exposed and shaded microhabitats are consistently associated with differences in physiological performance, and the frequency of occurrence of high temperatures is most probably limiting the maximum population densities supported at any given place. Topographical complexity provides thermal refugia throughout most of the distribution range, although towards the equatorial edges the magnitude of the amelioration provided by shaded microhabitats is largely reduced. Importantly, the limiting effects of temperature, rather than being related to latitude, seem to be tightly associated with microsite variability, which therefore is likely to have profound effects on the way local populations (and consequently species) respond to climatic changes.